The Strait of Gibraltar, the gateway between the Atlantic Ocean and the Mediterranean Sea, has a convulsive geological history, with recurring closing and opening events since the late Miocene. As a consequence, this region has played a major role in the evolutionary history of many species. Cynara baetica (Compositae) is a diploid perennial herb distributed in both sides of this strait. It is currently subdivided into two subspecies: C. baetica subsp. baetica for the Spanish populations, and C. baetica subsp. maroccana for the Moroccan ones. Following three different approximations of species delimitation, including phylogenetic and population genetic analyses (based on three AFLP primer combinations and two intergenic spacers of cpDNA), ecological niche modeling (ENM) and morphological studies, this taxon is investigated and reinterpreted. The results obtained showed a clear genetic, morphological and ecological differentiation between the two taxa and the important role played by the Strait of Gibraltar as a geographical barrier. Based on this evidence, the current taxonomic treatment is modified (both taxa should recover their specific rank) and specific conservation guidelines are proposed for the newly delimited taxa. ; An ADR fellowship (Universitat de Barcelona, Spain) was awarded to S. Massó for his PhD studies. This work has been supported by the Financial support from the Spanish Ministerio de Ciencia e Inovación (Spanish Government; projects CGL2007-60781 and CGL2010/18631) and the Agència de Gestió d'Ajuts Universitaris i de Recerca (Generalitat de Catalunya) (2009SGR0439 and 2014SGR514). ; Peer reviewed
9p., Diagramas. ; [EN] To assess temporal origins and diversification of lineages within subtribe Artemisiinae and Artemisia group a penalized likelihood analysis was applied on nrDNA ITS and ETS of 63 representatives. The tree was calibrated at the stem node of the Kaschgaria/ Artemisia lineage with the most reliable early Artemisia fossil pollen record from Late Oligocene (23 Ma). The results from this study suggest that the origin of the subtribe goes back to the Late Oligocene (24.6 ± 2.6 Ma) whilst the onset of differentiation of the genus Artemisia and most closely related genera is dated to the Early Miocene (19.8 ± 2.3 Ma). Divergence ages for lineages within the Artemisia group are often between the Early and Middle Miocene, whereas their radiations mostly occurred in the Late Miocene and Pliocene. The temporal context was also used to examine biogeographic and morphological (capitula and pollen type) evolution. Within the Artemisia group all lineages except the North American endemic have colonized the Mediterranean Basin at different epochs from Asian ancestors. Our analyses suggest the divergence of the North American endemic group from Asian ancestors (10.8 ± 1.5 Ma) in the Late Miocene. Homogamous-discoid capitula, characteristic of subgenera Seriphidium and Tridentatae, evolved not only in different geographic regions, but also at different times (2.0 ± 0.8 Ma and 7.9 ± 0.9 Ma respectively) within the Artemisia group. The loss of fertility of central flowers of disciform capitula should be considered as an ancient event in the genus since subgenus Dracunculus is one of the first groups that diverged (17.6 ± 2.1 Ma). ; [ES] Con el objetivo de estimar el tiempo de diversificación de los linajes de la subtribu Artemisiinae y el grupo Artemisia se aplicó un análisis de verosimilitud penalizado en base a las secuencias de ADNrn, ITS y ETS, de 63 representantes. El árbol se calibró en el nodo del linaje de Kaschgaria/Artemisia con el polen fósil de Artemisia que se consideró más fiablemente datado, siendo esta datación del Oligoceno Superior (23 Ma). Los resultados de este estudio datan el origen de la subtribu en el Oligoceno Superior (24,6 ± 2,6 Ma), mientras que la diferenciación del género Artemisia y los géneros relacionados es datada en el Mioceno Inferior (19,8 ± 2,3 Ma). El tiempo de divergencia de los linajes dentro del grupo Artemisia está situado entre el Mioceno Inferior y el Medio, mientras que la mayoría de sus radiaciones tuvieron lugar en el Mioceno Superior y en el Plioceno. El contexto temporal también se usó para examinar la evolución biogeográfica y morfológica (tipo de capítulo y polen). Todos los linajes dentro del grupo de Artemisia, excepto el complejo endémico norteamericano, han colonizado la cuenca mediterránea en diferentes épocas. Nuestros análisis sugieren la divergencia del grupo endémico norteamericano de antepasados asiáticos (10,8 ± 1,5 Ma) en el Mioceno Superior. El capítulo homógamo-discoide, característico de los subgéneros Seriphidium y Tridentatae, evolucionó no sólo en diferentes regiones geográficas, sino también en distintas épocas (2,0 ± 0,8 Ma y 7,9 ± 0,9 Ma respectivamente) dentro del grupo Artemisia. La pérdida de fertilidad de las flores centrales del capítulo disciforme se debería considerar como un evento antiguo en el género, puesto que el subgénero Dracunculus es uno de los primeros grupos que divergió (17,6 ± 2,1 Ma). ; This study was subsidized by projects CGL2004-04563-C02-02/ BOS of the Spanish government and the 2009/SGR/439 of the Generalitat de Catalunya. The first author received a predoctoral grant and support to perform a stay in the Botanical Garden of Vienna from the Spanish Government (FPI program). ; Peer reviewed
Genome size of 18 species of the genus Carduncellus, two species of the related genus Phonus and the monotypic genus Femeniasia (F. balearica) has been assessed by flow cytometry for the first time. Ploidy levels were assigned using genome size data together with previously reported chromosome counts. A phylogenetic framework was built to visualize how cytogenetic traits distributed across taxa. The results confirmed three ploidy levels (2x, 4x and 6x), with a predominance of diploids. The 2C values ranged from 3.24 pg in Carduncellus calvus to 11.16 pg in C. eriocephalus, whereas monoploid genome size (1Cx) ranged from 1.29 pg in C. duvauxii (4x) to 2.30 pg in Phonus rhiphaeus (2x). The mean 1Cx for tetraploids was lower than for diploids. For each ploidy level, genome size values of Carduncellus, Femeniasia and Phonus were found to be higher than those of Carthamus. This result is consistent with a trend frequently observed in plants, of higher genome sizes in long life cycle taxa compared to short-lived relatives. ; El tamaño del genoma de 18 especies del género Carduncellus, dos especies de los géneros relacionados, Phonus y el género monotípico Femeniasia (F. balearica) ha sido medido por primera vez mediante citometría de flujo. Los niveles de ploidía se asignaron utilizando datos de tamaño del genoma junto con los recuentos de cromosomas previamente reportados. Se construyó un marco filogenético para visualizar la distribución de las características citogenéticas de los táxones. Los resultados confirmaron tres niveles de ploidía (2x, 4x y 6x), con un predominio de los táxones diploides. Los valores de 2C oscilaron entre 3,24 pg en Carduncellus calvus y 11,16 pg en C. eriocephalus, mientras que el tamaño del genoma monoploide (1Cx) osciló entre 1,29 pg en C. duvauxii (4x) y 2,30 pg en Phonus rhiphaeus (2x). La media de los valores 1Cx para los tetraploides fue menor que para los diploides. Los valores de tamaño del genoma de Carduncellus, Femeniasia y Phonus fueron más elevados que los de Carthamus dentro del mismo nivel de ploidía. Este resultado concuerda con una tendencia frecuentemente observada en plantas en la que los táxones con ciclos de vida largos presentan tamaños del genoma más elevados que los táxones relacionados que poseen ciclos de vida cortos. ; This work has been supported by projects from the Spanish Government [(CGL2016-75694-P (AEI/FEDER, UE)] and the Generalitat de Catalunya (2017SGR1116). SG is the holder of a Ramón y Cajal contract (RYC-2014-16608) ; INTRODUCTION MATERIALS AND METHODS Plant material DNA content assessment Phylogenetic framework Statistical analyses RESULTS AND DISCUSSION CONCLUSIONS
19 p., mapas, tablas, ilustraciones ; The Macaronesian species of Euphorbia sect. Aphyllis subsect. Macaronesicae are distributed in four of the five archipelagos of Macaronesia and two mainland enclaves in Portugal and Morocco. The aims of this study are to investigate the biogeographic history of this group with AFLP and cpDNA markers, and to identify taxonomic entities within subsect. Macaronesicae based on genetic data, characterize them morphologically and infer the evolution of their diagnostic characters based on the reconstruction of ancestral character states. A continuous spatial diffusion analysis of AFLP data implicated Tenerife (central Canary Islands) as the area of origin of the group, followed by colonization of other Canarian islands and other Macaronesian archipelagos. Two dispersal events back to the mainland were also inferred. Our phylogenetic network, neighbour-joining clustering and Structure analyses of AFLP data demonstrated that species are genetically well delimited and suggested that they may have originated from a combination of allopatric speciation at broad scales (among islands) and fine scales (within islands), or possibly sympatric ecological speciation followed by more recent inter-island dispersal events. Ancestral character state reconstructions of morphological characters suggested that the ancestor of subsect. Macaronesicae was adapted to arid or mesic habitats, and traits associated with adaptation to humid habitats were acquired later. The central Canary Islands harbour the highest species diversity of this group in the Archipelago, and the highest nuclear and plastid genetic diversity. With regards to taxonomy, phylogenetic analyses and neighbour-joining clustering analyses based on AFLPs showed two clearly differentiated genetic groups, sister to each other, which correspond to the E. atropurpurea and E. lamarckii complexes formerly recognised based on morphology. Euphorbia aphylla is recovered as sister to the rest of the species, supporting its exclusion from the two complexes. Euphorbia tuckeyana is excluded from the E. lamarckii complex. ; This study was supported by the Ministry of Education and Science, Spanish Government (project CGL2010-18631/BOS, CGL2009-13322-C03-03, MEC-CSIC (200730i1035) and BES-2007-14260 (Ph.D. grant to L. Barres) and the Catalan Government ("Ajuts a grups consolidats" 2009/SGR/00439 and 2014/SGR/514). ; Peer reviewed
11 p. ; Sequences of the interna1 transcribed spacers (ITS1 and ITSZ) of nuclear ribosomal DNAwere analysed for 44Artemisia species (46 populations) representing al1 the five dassical subgenera and the geographical range of the genus, 11 species from 10 genera closely related to Artemisio, and six outgroup species from five other genera of the Anthemideae. The results definitely support the monophyly of the genus Artemisio in its broadest sense (including some taxa segregated as independent genera, like Oligosporus and Seriphidium). Eight main clades are established in this molecular phylogeny within Artemisio; they agree in part with the classical subdivision of the genus, but they also suggest that some infrageneric groups must be redefined. especially the subgenus Artemisio. The subgenera Tridentotae and ~eri~hidiuamre independent from each other. Some of the satellite G- enera are clearlv, ~. lacedw ithin Artemisio (Artemisiostrum, Filifolium, Mausoleo, Picrothomnus, Sphoeromerio, Turoniphytum), whereas some others fall outside the large clade formed by this genus (Brochonthemum, Elochanthemum, Hippolytio, Koschgorio). Our results, correlated to other data such as pollen morphology, allow us to conclude that the subtribe Artemisiinae as currently defined is a very heterogeneous group. Affinities of the largest genus of the subtribe and tribe,Artemisio, and of other genera of the subtribe to some genera from other subm'bes of the Anthemideae strongly suggest that subtribe Artemisiinae needs a deep revision and redefinition. Phylogenetic utility of region trnL-Fof the plastid DNA in the genus Artemisia and allies was also evaluated: sequences of the trnl-Fregion in Artemisio do not provide phylogenetic information. ; This work was subsidized by projects DGICYT PB–93–0032, DGICYT PB–97–1134 and BOS2001–3041–C02 of the Spanish government, and projects 1999SGR–00332 and 2001SGR–00125 of the Catalan government (through research group "Plant biodiversity and biosystematics"). ; Peer reviewed
16p., il. col., tablas y grafs. ; Euphorbia subg. Esula (Euphorbiaceae) has recently been shown, using molecular analyses, to contain a clade with a disjunct distribution in Macaronesia, South Africa and the Eritreo-Arabian region, and being primarily made up of members of sect. Tithymalus subsect. Pachycladae and sect. Tirucalli. To delimitate this disjoint group, we carried out phylogenetic analyses of the internal transcribed spacer (nrITS) using a broad sampling, with emphasis on subg. Esula. Subsequently, we carried out phylogenetic analyses focused on this clade using nuclear (ITS, ETS) and chloroplast (trnL-trnF, psbA-trnH, ycf3-trnS, trnG, atpB-rbcL, trnK-matK, trnT-trnL) markers, with the aim of resolving the phylogenetic relationships within the group and reconstructing its biogeographic history. Our results showed that sect. Tithymalus subsect. Pachycladae and sect. Tirucalli are polyphyletic. Section Aphyllis is recircumscribed to comprise the Pachycladae core clade and part of sect. Tirucalli. Low resolution within sect. Aphyllis and incongruences between nuclear and chloroplast phylogenies may be due to hybridization. Section Aphyllis should have originated in the Mediterranean area; its disjunct distribution is probably due to vicariance, resulting from fragmentation of a wider distribution area in North Africa caused by the aridification of the climate during the late Miocene-Pliocene. ; This work was supported by the Ministry of Education and Science, Spanish Government (project CGL2006-01765/BOS, CGL2009- 13322-C03-03 and BES-2007-14260 Ph.D. grant to L. Barres) and the Catalan Government ('Ajuts a grups consolidats' 2009/SGR/00439). We acknowledge the Euphorbia Planetary Biodiversity Inventory Project, which provided material and strongly supports our research. Special thanks are due to A. Moller and J. Morawetz, who kindly provided some photographs for Figure 3. The National Science Foundation Planetary Biodiversity Inventory Grant (DEB-0616533) additionally supported this work ; Peer reviewed
25 p., figuras, tablas -- Postprint del artículo publicado en Taxon. Versión revisada y corregida -- Contiene material suplementario ; Multiple factors related to complex geomorphological and climatic history, in addition to biological factors such as hybridization, hinder the definition of some Mediterranean plant groups. The existence of controversial taxonomic treatments, the possible hybridization events involved, and its unknown evolutionary history, make the Helichrysum pendulum complex of H. sect. Stoechadina an ideal model to understand general processes of Mediterranean plant systematics and evolution. The mosaic range of the complex, which is distributed over several islands and continental areas in the western-central Mediterranean Basin, provides an opportunity to investigate how past connections and disconnections between landmasses may have determined the current geographic distribution of genetic variation in this area. The cpDNA region rpl32-trnL intergenic spacer and the nrDNA region ETS were sequenced for 1–8 individuals from each of the 44 populations sampled, covering all taxa and the whole geographic range of the complex. These individuals were analysed together with a broad sampling of the remaining members of sect. Stoechadina. In addition, detailed multivariate analyses of morphological characters were performed for the whole section and for the H. pendulum complex. Considering together distinctive genetic and morphological traits, our species concept is presented and discussed in a context of integrative taxonomy, and five species are recognized within the complex: H. errerae, H. melitense, H. pendulum, H. saxatile and H. valentinum. The first three species are recognizable by qualitative and quantitative morphological traits, and are genetically distinguishable from the rest as shown by the molecular markers analysed. The two last species are reported here to have a putative ancient hybrid origin and are also genetically distinguishable from the rest but morphologically recognisable only by quantitative characters. Phylogenetic relationships shown by nuclear and chloroplast markers, and an intermediate morphology between the two putative parental taxa, point to H. pendulum and H. italicum as the putative parental taxa for H. saxatile, and H. pendulum and H. stoechas as putative parental taxa for H. valentinum. In a discriminant analysis of the five species, 97.8% of all individuals were classified correctly. The high level of haplotype and ribotype diversity observed in North Africa indicates that this region is either the area of origin of the complex or a secondary contact zone. Our results suggest that the complex colonized several islands and migrated through the Gibraltar and Sicilian Straits during phases of low sea level, favoured by local dispersal events that promoted its gradual range expansion. The occurrence of the complex in the Balearic Islands, which have remained isolated even during low sea level phases, could be explained by stochastic long-distance dispersal events. ; Financial support from the Spanish Ministerio de Ciencia e Inovación CGL2007-60781/BOS, CGL2009-13322-C03-03/BOS, CGL2010-18631/BOS) and the Catalan government ('Ajuts a grups consolidats' 2009/SGR/00439 and 2014/SGR/514) is also acknowledged. ; Peer reviewed
16 p. ; Premise of the study: Tribe Cardueae (thistles) forms one of the largest tribes in the family Compositae (2400 species), with representatives in almost every continent. The greatest species richness of Cardueae occurs in the Mediterranean region where it forms an important element of its flora. New fossil evidence and a nearly resolved phylogeny of Cardueae are used here to reconstruct the spatiotemporal evolution of this group. Methods: We performed maximum parsimony and Bayesian phylogenetic inference based on nuclear ribosomal DNA and chloroplast DNA markers. Divergence times and ancestral area reconstructions for main lineages were estimated using penalized likelihood and dispersal–vicariance analyses, respectively, and integrated over the posterior distribution of the phylogeny from the Bayesian Markov chain Monte Carlo analysis to accommodate uncertainty in phylogenetic relationships. Key results: The phylogeny shows that subtribe Cardopatiinae is sister to the remaining subtribes, and subtribes Carlininae and Echinopsinae appear as consecutive sister-clades to the Carduinae/Centaureinae. Tribe Cardueae is inferred to have originated around the Mid Eocene in West Asia, which is also the ancestral area of most subtribes within Cardueae. Diversification within each subtribe began during the Oligocene-Miocene period. Conclusions: Most diversification events within Cardueae are related to the continuous cycles of area connection and division between the Anatolian microplate and the western Mediterranean Basin during the Oligocene-Miocene and with the uplift of the Himalayan range from the Miocene onward. From these two regions, thistles dispersed and colonized the rest of the continents (e.g., the New World, Africa, and Australia), most likely during the colder Pliocene-Pleistocene period. ; The authors thank the Ministry of Education and Science of the Spanish Government (projects CGL2006-01765/BOS and CGL2009-13322-C03-03/BOS, CGL2009-13322-C03-01/BOS, CGL2010-18631 and grant BES-2007-14260 to L.B.) and the Catalan Government ('Ajuts a grups consolidats' 2009/SGR/00439) for financial support. S.B. and C.L.A. were supported by postdoctoral grants at RJB from the Swiss National Science Foundation grant (PBNEP3-129903) and the Swedish Research Council, respectively. ; Peer reviewed
